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Large-scale habitat segregation of fish-eating and mammal-eating killer whales (Orcinus orca) in the western North Pacific

Polar Biology volume 42, pages 931–941 (2019)Cite this article

Abstract

Top predators strongly impact the structure of ecosystems through the top-down cascading effects on prey species. Killer whales (Orcinus orca), the top predators in marine ecosystems, are increasing their presence in the Arctic following the ice cover loss. The impact of killer whales on marine ecosystems differs dramatically across ecotypes: ‘resident’ R-type killer whales feed mostly on large fish, while ‘transient’ T-type whales feed mostly on marine mammals. We analyze the differences in geographical distribution of killer whale ecotypes in the western North Pacific in order to predict the potential effects of climate change on their range and the consequential impact on Arctic ecosystems. We show that R-type whales prevail in the coastal waters of eastern Kamchatka, Commander and Kuril Islands and in the central Okhotsk Sea, while T-type whales dominate the coastal waters of Chukotka and the coastal Okhotsk Sea. The most prominent difference between these areas is depth: in the regions where R-type whales prevail, deep waters occur close to shore, while the regions dominated by T-type whales are represented by wide shallows covered with ice in winter. We propose several hypotheses to explain this large-scale segregation, including distribution of prey species and ice cover. Habitat preferences suggest that range expansion in the Arctic induced by climate change will likely involve mostly T-type killer whales and consequently increase the predation pressure on marine mammals to a greater extent than on fish stocks. This should be considered in further studies and prediction models highlighting Arctic marine ecosystems change.

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References

  • Baird RW, Hanson MB, Dill LM (2005) Factors influencing the diving behaviour of fish-eating killer whales: sex differences and diel and interannual variation in diving rates. Can J Zool 83:257–267

    Article  Google Scholar 

  • Baird RW, Dill LM (1995) Occurrence and behaviour of transient killer whales: seasonal and pod-specific variability, foraging behaviour, and prey handling. Can J Zool 73:1300–1311

    Article  Google Scholar 

  • Baird RW, Dill LM (1996) Ecological and social determinants of group size in transient killer whales. Behav Ecol 7:408–416

    Article  Google Scholar 

  • Baird RW, Stacey PJ (1988) Variation in saddle patch pigmentation in populations of killer whales (Orcinus orca) from British Columbia, Alaska, and Washington State. Can J Zool 66:2582–2585

    Article  Google Scholar 

  • Barrett-Lennard LG, Ford JK, Heise KA (1996) The mixed blessing of echolocation: differences in sonar use by fish-eating and mammal-eating killer whales. Animal Behav 51:553–565

    Article  Google Scholar 

  • Bigg MA (1982) An assessment of killer whale (Orcinus orca) stocks off Vancouver Island, British Columbia. Rep Int Whaling Comm 32:655–666

    Google Scholar 

  • Bjørge A (2001) How persistent are marine mammal habitats in an ocean of variability? In: Evans PGH, Raga JA (eds) Marine mammals: biology and conservation. Kluwer Academic/Plenum Publishers, New York, pp 63–91

    Google Scholar 

  • Breed GA, Matthews CJ, Marcoux M, Higdon JW, LeBlanc B, Petersen SD, Orr J, Reinhart NR, Ferguson SH (2017) Sustained disruption of narwhal habitat use and behavior in the presence of Arctic killer whales. Proc Natl Acad Sci USA 114:2628–2633

    Article  CAS  PubMed  Google Scholar 

  • Burdin AM, Hoyt E, Sato H, Tarasyan KK, Filatova OA (2005) Resident and transient-type Killer whales, Orcinus orca, in Southeast Kamchatka, Russia. Int Whal Comm Rep SC/56/SM15

  • Burt WH (1943) Territoriality and home range concepts as applied to mammals. J Mamm 24:346–352

    Article  Google Scholar 

  • Deecke VB, Ford JK, Slater PJ (2005) The vocal behaviour of mammal-eating killer whales: communicating with costly calls. Animal Behav 69:395–405

    Article  Google Scholar 

  • Emmons CK, Hard JJ, Dahlheim ME, Waite JM (2018) Quantifying variation in killer whale (Orcinus orca) morphology using elliptical Fourier analysis. Mar Mam Sci. https://doi.org/10.1111/mms.12505

    Article  Google Scholar 

  • Eschmeyer WN, Herald ES, Hammann H (1983) A field guide to Pacific coast fishes of North America. Houghton Mifflin Company, Boston

    Google Scholar 

  • Ferguson SH, Higdon JW, Chmelnitsky EG (2010) The rise of killer whales as a major Arctic predator. In: Ferguson SH, Loseto LL, Mallory ML (eds) A little less Arctic. Springer, Dordrecht, pp 117–136

    Chapter  Google Scholar 

  • Filatova OA, Borisova EA, Shpak OV, Meschersky IG, Tiunov AV, Goncharov AA, Burdin AM (2015) Reproductively isolated ecotypes of killer whales Orcinus orca in the seas of the Russian Far East. Biol Bull 42:674–681

    Article  Google Scholar 

  • Filatova OA, Borisova EA, Meschersky IG, Logacheva MD, Shpak OV, Hoyt E (2018) Colonizing the wild west: low diversity of complete mitochondrial genomes in western North Pacific resident killer whales suggests the founder effect. J Hered 109:735–743

    PubMed  Google Scholar 

  • Ford JKB, Ellis GM, Barrett-Lennard LG, Morton AB, Palm RS, Balcomb KC (1998) Dietary specialization in two sympatric populations of killer whales (Orcinus orca) in coastal British Columbia and adjacent waters. Can J Zool 76:1456–1471

    Article  Google Scholar 

  • Ford JKB, Ellis GM, Balcomb KC (2000) Killer whales: the natural history and genealogy of Orcinus orca in British Columbia and Washington. UBC Press, Vancouver

    Google Scholar 

  • Ford JKB, Ellis GM, Matkin CO, Wetklo MH, Barrett-Lennard LG, Withler RE (2011) Shark predation and tooth wear in a population of northeastern Pacific killer whales. Aquat Biol 11:213–224

    Article  Google Scholar 

  • Ford JK, Ellis GM (2006) Selective foraging by fish-eating killer whales Orcinus orca in British Columbia. Mar Ecol Prog Ser 316:185–199

    Article  Google Scholar 

  • Ford JKB, EH Stredulinsky, Towers JR, Ellis GM (2013) Information in support of the identification of critical habitat for transient killer whales (Orcinus orca) off the West Coast of Canada. DFO Can Sci Advis Sec Res Doc 2012/155, p 46

  • Fossheim M, Primicerio R, Johannesen E, Ingvaldsen RB, Aschan MM, Dolgov AV (2015) Recent warming leads to a rapid borealization of fish communities in the Arctic. Nat Clim Chang 5:673–677

    Article  Google Scholar 

  • Frost KJ, Russell RB, Lowry LF (1992) Killer whales, Orcinus orca, in the southeastern Bering Sea: Recent sightings and predation on other marine mammals. Mar Mamm Sci 8:110–119

    Article  Google Scholar 

  • Gorbarenko SA, Psheneva OY, Artemova AV, Tiedemann R, Nürnberg D (2010) Paleoenvironment changes in the NW Okhotsk Sea for the last 18 kyr determined with micropaleontological, geochemical, and lithological data. Deep Sea Res Part I 57:797–811

    Article  CAS  Google Scholar 

  • Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symp Ser 41:95–98

    CAS  Google Scholar 

  • Hauser DDW (2006) Summer space use of Southern resident killer whales (Orcinus orca) within Washington and British Columbia inshore waters. Master thesis, University of Washington, Seattle

  • Heimlich-Boran JR (1988) Behavioral ecology of killer whales (Orcinus orca) in the Pacific Northwest. Can J Zool 66:565–578

    Article  Google Scholar 

  • Higdon JW, Ferguson SH (2009) Loss of Arctic sea ice causing punctuated change in sightings of killer whales (Orcinus orca) over the past century. Ecol Appl 19:1365–1375

    Article  PubMed  Google Scholar 

  • Hoelzel AR, Dahleim M, Stern SJ (1998) Low genetic variation among killer whales (Orcinus orca) in the Eastern North Pacific and genetic differentiation between foraging specialists. J Hered 89:121–128

    Article  CAS  PubMed  Google Scholar 

  • Hoelzel AR, Goldsworthy SD, Fleischer RC (2002) Population genetics. In: Hoelzel AR (ed) Marine mammal biology: an evolutionary approach. Blackwell Science, Oxford, pp 325–352

    Google Scholar 

  • Ivkovich TV, Filatova OA, Burdin AM, Sato H, Hoyt E (2010) The social organization of resident-type killer whales (Orcinus orca) in Avacha Gulf, Northwest Pacific, as revealed through association patterns and acoustic similarity. Mamm Biol 75:198–210

    Article  Google Scholar 

  • Jay CV, Fischbach AS, Kochnev AA (2012) Walrus areas of use in the Chukchi Sea during sparse sea ice cover. Mar Ecol Prog Ser 468:1–13

    Article  Google Scholar 

  • Kornev SI, Belonovich OA, Nikulin VS (2014) Killer whales (Orcinus orca) and Greenland halibut (Reinhardtius hippoglossoides) fishery in the Sea of Okhotsk. Res Water Biol Resour Kamchatka N West Part Pac Ocean 34:35–50

    Google Scholar 

  • Kortsch S, Primicerio R, Fossheim M, Dolgov AV, Aschan M (2015) Climate change alters the structure of arctic marine food webs due to poleward shifts of boreal generalists. Proc R Soc B 282:20151546

    Article  PubMed  Google Scholar 

  • Laidre KL, Stern H, Kovacs KM, Lowry L, Moore SE, Regehr EV, Ferguson SH, Wiig Ø, Boveng P, Angliss RP, Born EW, Litovka D, Quakenbush L, Lydersen C, Vongraven D, Ugarte F (2015) Arctic marine mammal population status, sea ice habitat loss, and conservation recommendations for the 21st century. Conserv Biol 29:724–737

    Article  PubMed  PubMed Central  Google Scholar 

  • Laidre KL, Regehr EV (2017) Arctic marine mammals and sea ice. In: Thomas DN (ed) Sea ice, 3rd edn. Wiley, New York, pp 516–533

    Google Scholar 

  • MacCracken JG (2012) Pacific walrus and climate change: observations and predictions. Ecol Evol 2:2072–2090

    Article  PubMed  PubMed Central  Google Scholar 

  • Matthews CJ, Luque SP, Petersen SD, Andrews RD, Ferguson SH (2011) Satellite tracking of a killer whale (Orcinus orca) in the eastern Canadian Arctic documents ice avoidance and rapid, long-distance movement into the North Atlantic. Polar Biol 34:1091–1096

    Article  Google Scholar 

  • Melnikov VV, Zagrebin IA (2005) Killer whale predation in coastal waters of the Chukotka Peninsula. Mar Mamm Sci 21:550–556

    Article  Google Scholar 

  • Miller PJO, Shapiro AD, Deecke VB (2010) The diving behaviour of mammal-eating killer whales (Orcinus orca): variations with ecological not physiological factors. Can J Zool 88:1103–1112

    Article  Google Scholar 

  • Morin PA, Archer FI, FooteAD Vilstrup J, Allen EE, Wade P, Harkins T (2010) Complete mitchondrial genome phylogeographic analysis of killer whales (Orcinus orca) indicates multiple species. Genome Res 20:908–916

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Morton AB (1990) A quantitative comparison of the behaviour of resident and transient forms of the killer whale off the central British Columbia coast. Rep Int Whal Comm Special Issue 12:245–248

    Google Scholar 

  • Nichol LM, Shackleton DM (1996) Seasonal moving and foraging behavior of northern resident killer whales (Orcinus orca) relation to inshore distribution of salmon (Oncorhynchus spp.) in British Columbia. Can J Zool 74:983–991

    Article  Google Scholar 

  • Niebauer HJ, Schell DM (1993) Physical environment of the Bering Sea population. In: Burns JJ, Montague JJ, Cowles CJ (eds) The bowhead whale, 1st edn. Society for Marine Mammalogy, Lawrence, pp 23–43

    Google Scholar 

  • Nosil P (2012) Ecological speciation. Oxford University Press, Oxford

    Book  Google Scholar 

  • O'Corry-Crowe G, Suydam RS, Rosenberg A, Frost KJ, Dizon AE (1997) Phylogeography, population structure and dispersal patterns of the beluga whale Delphinapterus leucas in the western Nearctic revealed by mitochondrial DNA. Mol Ecol 6:955–970

    Article  CAS  Google Scholar 

  • Pante E, Simon-Bouhet B (2013) marmap: a package for importing, plotting and analyzing bathymetric and topographic data in R. PLoS ONE 8:e73051

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Parsons KM, Durban JW, Burdin AM, Burkanov VN, Pitman RL, Barlow J, Wade PR (2013) Geographic patterns of genetic differentiation among killer whales in the northern North Pacific. J Hered 104:737–754

    Article  PubMed  Google Scholar 

  • Pitman RL, Ensor P (2003) Three forms of killer whales (Orcinus orca) in Antarctic waters. J Cet Res Manag 5:131–140

    Google Scholar 

  • R Core Team (2014) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.https://www.R-project.org/

  • The IUCN Red List of Threatened Species. Version 2017-3. www.iucnredlist.org. Accessed 31 May 2018

  • Reeves RR, Perrin WF, Taylor BL, Baker CS, Mesnick SL (2004) Report of the workshop on shortcomings of cetacean taxonomy in relation to needs of conservation and management, La Jolla California. NOAA Technical Memorandum NMFS SWFSC 363

  • Rice A, Deecke VB, Ford JK, Pilkington JF, Oleson EM, Hildebrand JA (2017) Spatial and temporal occurrence of killer whale ecotypes off the outer coast of Washington State, USA. Mar Ecol Prog Ser 572:255–268

    Article  Google Scholar 

  • Saulitis E, Matkin C, Barrett-Lennard L, Heise K, Ellis G (2000) Foraging strategies of sympatric killer whale (Orcinus orca) populations in Prince William Sound, Alaska. Mar Mamm Sci 16:94–109

    Article  Google Scholar 

  • Saulitis E, Holmes LA, Matkin C, Wynne K, Ellifrit D, St-Amand C (2015) Biggs killer whale (Orcinus orca) predation on subadult humpback whales (Megaptera novaeangliae) in Lower Cook Inlet and Kodiak. Alaska. Aquat Mamm 41:341

    Article  Google Scholar 

  • Scheel D, Matkin CO, Saulitis E (2001) Distribution of killer whale pods in Prince William sound, Alaska 1984–1996. Mar Mamm Sci 17:555–569

    Article  Google Scholar 

  • Schluter D (2000) The ecology of adaptive radiation. Oxford Univ Press, Oxford

    Google Scholar 

  • Shelden KE, Rugh DJ, Mahoney B, Dahlheim ME (2003) Killer whale predation on belugas in Cook Inlet, Alaska: implications for a depleted population. Mar Mamm Sci 19:529–544

    Article  Google Scholar 

  • Shoener TW (1974) Resource partitioning in ecological communities. Science 185:27–39

    Article  Google Scholar 

  • Shpak OV, Paramonov AYu (2018) The bowhead whale, Balaena mysticetus Linnaeus, 1758, in the western Sea of Okhotsk (2009–2016): distribution pattern, behavior, and threats. Russ J Mar Biol 44:210–218

    Article  CAS  Google Scholar 

  • Shpak OV, Shulezhko TS (2013) Observations and photoidentification of an unusual group of mammal-eating killer whales (Orcinus orca) in the western Sea of Okhotsk. Res Water Biol Resour Kamchatka N West Part Pac Ocean 28:129–139

    Google Scholar 

  • Shulezhko TS, Permyakov PA, Ryazanov SD, Burkanov VN (2018) Bigg's Killer Whales (Orcinus orca) in the Kuril Islands. Aquat Mamm 44:267–278

    Article  Google Scholar 

  • Shuntov VP (2008) Pacific salmons in marine and oceanic ecosystems. TINRO-Centr, Vladivostok

    Google Scholar 

  • Silber GK, Lettrich M, Thomas PO, Baker J, Baumgartner M, Becker EA, Boveng P, Dick D, Fiechter J, Forcada J, Forney K, Griffis R, Hare J, Hobday A, Howell D, Laidre KL, Mantua N, Quakenbush L, Santora JA, Stafford KM, Spencer P, Stock C, Sydeman WJ, Van Houtan K, Waples RS (2017) Projecting marine mammal distribution in a changing climate. Front Mar Sci 4:413

    Article  Google Scholar 

  • Solovyev BA, Shpak OV, Glazov DM, Rozhnov VV, Kuznetsova DM (2015) Summer distribution of beluga whales (Delphinapterus leucas) in the Sea of Okhotsk. Russ J Theriol 14:201–215

    Article  Google Scholar 

  • Solovyeva M, Glazov D, Kuznetsova D, Burkanov V, Boveng P, London J. Rozhnov V (2017) Seals of the Okhotsk Sea: migrations and ecology as context for comparison with Bering Sea populations. Book of Abstracts, Alaska Marine Science Symposium, p 350

  • Tarasyan KK, Filatova OA, Burdin AM, Hoyt E, Sato H (2005) Keys for the status of killer whales in Eastern Kamchatka, Russia: foraging ecology and acoustic behaviour. Biosphere Conser 6:73–83

    Google Scholar 

  • Towers JR, Tixier P, Ross KA, Bennett J, Arnould JPY, Pitman RL, Durban JW (2018) Movements and dive behaviour of a toothfish-depredating killer and sperm whale. ICES J Mar Sci. https://doi.org/10.1093/icesjms/fsy118

    Article  Google Scholar 

  • Volkova EV, Ivkovich TV, Shitova MV, Chernyaeva E, Malinina TV, Okorokova SS, Burdin AM, Hoyt E (in rev) The summer diet of fish-eating killer whales in Avacha Gulf of Kamchatka: are there any preferences? Mamm Biol

  • Wassmann P, Duarte CM, Agusti S, Sejr MK (2011) Footprints of climate change in the Arctic marine ecosystem. Glob Change Biol 17:1235–1249

    Article  Google Scholar 

  • Wright BM, Ford JKB, Ellis GM, Deecke VB, Shapiro AD, Battaile BC, Trites AW (2017) Fine-scale foraging movements by fish-eating killer whales (Orcinus orca) relate to the vertical distributions and escape responses of salmonid prey (Oncorhynchus spp.). Mov Ecol 5:3

  • Zerbini AN, Waite JM, Durban JW, LeDuc R, Dahlheim ME, Wade PR (2007) Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling. Mar Biol 150:1033–1045

    Article  Google Scholar 

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Acknowledgements

This research was supported by the Pew Marine Fellowship for OF and the Russian Fund for Fundamental Research (Grant 18–04-00462). The expeditions that provided the data for this study were funded by a variety of sources including Whale and Dolphin Conservation, Animal Welfare Institute, Humane Society International, the Litowitz Foundation, Alaska SeaLife Center, Russian Geographical Society, Pew Marine Fellowship, Russian Fund for Fundamental Research. We are grateful to all the people who kindly shared the photographs of their opportunistic encounters with killer whales including Alexandr Bobkov, Elena Golubova, Sergey Gorshkov, Vladimir Dekin, Sergey Fomin, Karina Karenina, Mikhail Korostelev, Maxim Kozlov, Vyacheslav Kozlov, Evgeny Lebedev, Viktor Nikulin, Sergey Nikulin, Nikolay Pavlov, Lika Pokrovskaya, Alexey Timshin, Grigory Tsidulko, Jean-Pierre Sylvestre, Ivan Usatov, Peter Van der Wolf, Vladimir Vertuankin, and David Weller. We also thank all the students and volunteers who participated in our expeditions and helped to collect the data presented here.

We are grateful to Jared Towers and an anonymous reviewer for their thoughtful comments and corrections of the manuscript.

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Authors and Affiliations

  1. Faculty of Biology, Moscow State University, Moscow, Russia, 119234

    Olga A. Filatova & Ivan D. Fedutin

  2. Severtsov Institute of Ecology and Evolution RAS, Moscow, Russia, 119071

    Olga V. Shpak

  3. Faculty of Biology, St. Petersburg State University, St. Petersburg, Russia, 199034

    Tatiana V. Ivkovich & Evgeniia V. Volkova

  4. School of Veterinary Science, The University of Queensland, Brisbane, Australia

    Ekaterina N. Ovsyanikova

  5. Kamchatka Branch of Pacific Institute of Geography FEB RAS, Petropavlovsk-Kamchatsky, Russia, 683024

    Alexander M. Burdin

  6. Whale and Dolphin Conservation, Bridport, DT6 5DD, Dorset, UK

    Erich Hoyt

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  1. Olga A. Filatova
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  2. Olga V. Shpak
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  3. Tatiana V. Ivkovich
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  4. Evgeniia V. Volkova
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  7. Alexander M. Burdin
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  8. Erich Hoyt
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Filatova, O.A., Shpak, O.V., Ivkovich, T.V. et al. Large-scale habitat segregation of fish-eating and mammal-eating killer whales (Orcinus orca) in the western North Pacific. Polar Biol 42, 931–941 (2019). https://doi.org/10.1007/s00300-019-02484-6

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Keywords

  • Habitat segregation
  • Killer whale
  • Ecotypes
  • Climate change
  • Arctic